Beyond “all-or-nothing” climbing fibers: graded representation of teaching signals in Purkinje cells

Farzaneh Najafi, Javier F. Medina

Arguments about the function of the climbing fiber (CF) input to the cerebellar cortex have fueled a rabid debate that started over 40 years ago, and continues to polarize the field to this day. The origin of the controversy can be traced back to 1969, the year David Marr published part of his dissertation work in a paper entitled "A theory of cerebellar cortex." In Marr's theory, CFs play a key role during the process of motor learning, providing an instructive signal that serves as a "teacher" for the post-synaptic Purkinje cells. Although this influential idea has found its way into the mainstream, a number of objections have been raised. For example, several investigators have pointed out that the seemingly "all-or-nothing" activation of the CF input provides little information and is too ambiguous to serve as an effective instructive signal. Here, we take a fresh look at these arguments in light of new evidence about the peculiar physiology of CFs. Based on recent findings we propose that at the level of an individual Purkinje cell, a graded instructive signal can be effectively encoded via pre- or post-synaptic modulation of its one and only CF input.